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Abstract

An endemic genus of Madagascan spiders (Araneae, Archaeidae, Eriauchenius) is revised. All 20 species of Eriauchenius are described and keyed, of which 14 are new species: Eriaucheniusandriamanelosp. n., Eriaucheniusandrianampoinimerinasp. n., Eriaucheniusgoodmanisp. n., Eriaucheniusharveyisp. n., Eriaucheniuslukemacaulayisp. n., Eriaucheniusmilajaneaesp. n., Eriaucheniusmillotisp. n., Eriaucheniusrafohysp. n., Eriaucheniusranavalonasp. n., Eriaucheniusrangitasp. n., Eriaucheniusrixisp. n., Eriaucheniussamasp. n., Eriaucheniuswunderlichisp. n., Eriaucheniuszirafysp. n. Additionally, six species of the new genus Madagascarchaeagen. n. are described and keyed, of which four are new species: Madagascarchaeafohysp. n., Madagascarchaealotzisp. n., Madagascarchaeamoramorasp. n., Madagascarchaearabesahalasp. n. Diagnostic characters for the Madagascan and African genera are described, and based on these characters and previous phylogenetic analyses the following species transfers are proposed: Eriaucheniuscornutus (Lotz, 2003) to Afrarchaea; Afrarchaeafisheri (Lotz, 2003) and Afrarchaeamahariraensis (Lotz, 2003) to Eriauchenius. Finally, we propose that the distribution of Afrarchaea be restricted to South Africa. While several Madagascan specimens have previously been identified as Afrarchaeagodfreyi (Hewitt, 1919), we argue that these are likely misidentifications that should instead be Eriauchenius.

Keywords

Afrarchaea, Palpimanoidea, new species, taxonomy

Introduction

Archaeid spiders, commonly called pelican or assassin spiders, are an ancient, paleoendemic group that has existed since Pangaean times (Wood et al. 2013). These spiders do not build a web to capture their prey. Instead, they are active hunters and they have an unusual modification to their cephalic area that relates to their predatory behaviors: the carapace is extended and tubular in structure, and encircles the cheliceral bases, giving archaeids the appearance of a ‘‘neck’’ and “head”; the chelicerae are also greatly elongated. While most spiders are predatory generalists (Foelix 2011), archaeids are highly specialized and will only prey on other spiders (Legendre 1961; Millot 1948; Wood 2008; Wood et al. 2012). The highly modified cephalic area in archaeids is used to employ a novel prey capture strategy that is unique among spiders (see fig. 1 in Wood et al. 2007): the modified carapace allows for highly maneuverable and elongated chelicerae that can be extended 90° away from the body to attack spider prey at a distance (Forster and Platnick 1984; Wood et al. 2012). There is a diversity of carapace shapes among archaeids, with ‘‘necks’’ of varying degrees of elongation, from long and constricted to short and stout. Reliance on the elevated cephalic area as a phylogenetically informative character has served as the basis for historical classifications of archaeid spiders and their closest relatives (Forster and Platnick 1984; Legendre 1970; Platnick 1991). However, it has been shown that different morphs with varying degrees of elevation in the cephalic area have evolved in parallel within the family (Wood et al. 2015; Wood et al. 2007).

The morphological interspecific diversity in the carapace and chelicerae shape seems to relate to their global distribution and diversification patterns. Archaeid spiders have distinct Northern Hemisphere lineages, now extinct and known only from fossils dated from the Eocene to the Jurassic (Koch and Berendt 1854; Penney 2003; Selden et al. 2008; Wood 2017; Wunderlich 2004, 2008, 2015). The extant lineages occur only in the Southern Hemisphere, restricted to Madagascar, mainland Australia, and South Africa (World Spider Catalog 2017). A divergence dating study that included both fossil and extant taxa concluded that the Southern Hemisphere lineages are monophyletic and that the split between the extinct northern and extant southern faunas likely relates to Pangaea breaking into Gondwana and Laurasia in the Jurassic (Wood et al. 2013). So, it appears that archaeids were once more widespread, but are currently restricted to relictual areas. Furthermore, there has been a shift in morphological features through time: the fossil archaeids in general had shorter carapace/chelicerae features, and occupied a unique region of archaeid morphospace, whereas the extant clades have more elongated features (see fig. 5 in Wood 2017). While the northern lineages went extinct, the southern lineages persisted, and within their present day relictual distributions the extant clades have diversified. Recent studies have recovered the timing of diversification in the extant Australian clade to be congruent with Miocene aridification (Rix and Harvey 2012c), and in the South African clade to be congruent with Miocene uplift of the Great Escarpment (Wood et al. 2015). Madagascan lineages have been exposed to more ancient and more turbulant geoclimatic events, and compared to the Australian and South African clades, the Madagascan clades show an increased rate of morphological evolution in carapace and chelicerae shape (Wood et al. 2015). Furthermore, there is a greater degree of sympatry among Madagascan species than is observed in the Australian and South African species. Morphological diversification in the Madagascan clade may be due to exposure to more ancient geoclimatic events that led to the build-up of sympatric species in montane rainforest areas (Wood et al. 2015).

Phylogenetic analysis of molecular and morphological data by Wood et al. (2012) placed archaeids in the superfamily Palpimanoidea along with four other families. However, recent phylogenetic analyses of molecular data failed to recover this same Palpimanoidea clade, although, the monophyly of Archaeidae is strongly supported and not disputed (Wheeler et al. 2016). Since Forster and Platnick (1984) delimited the family to 17 species, there have been subsequent revisions that have expanded our knowledge of archaeid species diversity: the Australian archaeids are nearly completely revised (Rix and Harvey 2011, 2012a, b) for a total of 38 species; 12 South African species have been described (Lotz 1996, 2003, 2006, 2015), although new species are still being discovered (Wood et al. 2015); the “gracilicollis group” from Madagascar was revised (Wood 2008) and contains 14 species; and here, we perform a revision of the Madagascan Eriauchenius. A recent phylogenetic analysis (Wood et al. 2015) resolved the relationships among the extant genera with strong branch support: the Australian lineages were found to be monophyletic and sister to the African + Madagascar species; there are two monophyletic groups on Madagascar, Eriauchenius and Madagascarchaea gen. n.; the South African clade Afrarchaea Forster & Platnick, 1984 is sister to Eriauchenius, and Afrarchaea + Eriauchenius is sister to Madagascarchaea gen. n. The phylogeny from Wood et al (2015) is reproduced here, but edited to include the names of the new genus and the newly described species (Fig. 1). Based on this phylogeny as well as the morphological examination of species, in the current study we propose several taxonomic changes: first, the creation of a new genus Madagascarchaea gen. n., which was previously revised as the “gracilicollis group” (Wood 2008); and second, the transfer of several species to different genera. The current paper also performs a taxonomic revision of Eriauchenius, describing 14 new species, and additionally 4 new species of Madagascarchaea gen. n.

Figure 1.

Total evidence phylogeny from Bayesian analysis of molecular and morphological data, from Wood et al. (2015). Outgroups not shown and with the morphospecies names changed to the new species names from the current study. Posterior probabilities are as follows, greater than or equal to: (*) 0.90; (**) 0.95; (***) 0.99.

Methods

Specimens examined in this study were primarily from the California Academy of Sciences collection. Additional material was borrowed from the museums referred to in Table 1, which lists the museum abbreviations used in the text. Scanning electron microscope (SEM) images were taken using a Leo 1450VP, a JEOL JSM-6335F, and a Zeiss EVO MA15 scanning electron microscope; prior to photographing, specimens were critically point dried and sputter coated. Specimens were also examined and interpretively drawn using a Leica MZ16 and M205C microscope that had a drawing tube attached. The vulva, after dissection, was placed overnight in Ultrazyme contact lens cleaner. Photographs were taken as a series of stacks using a Nikon DXM1200 or a Canon EOS T6i digital camera mounted on the Leica microscope or using a BK Plus Laboratory System from Visionary Digital (Palmyra, PA, USA) equipped with a Canon EOS 7D camera. Image stacks were assembled using stacking software Auto-Montage Pro (Syncroscopy, Digital Imaging Systems Ltd.), HeliconFocus (Helicon Soft Ltd.) or ZereneStacker (Zerene Systems, LLC). All measurements are in millimetres (mm). Morphological abbreviations used in the text are listed in Table 2. The pars cephalica in Archaeidae is elongated and the term ‘head’ or ‘cephalon’ represents the most distal portion of the elongation, and the term ‘neck’ represents the constricted portion of the elongation. The measurement CtH is the length from the lateral edge of the carapace between coxae I and II to the top of the cephalon, running parallel with the “neck” (Fig. 2). This measurement takes the tilt of the ‘neck’ into account. The carapace tilt angle is defined as the angle between the posterior edge of the lateral side of the carapace (the portion above coxae II and III) and the anterior edge of the “neck” in the lateral view (Fig. 2). The CtH/CL ratio quantifies the elongation of the cephalic area by dividing the CtH by the carapace length. The female genitalia have a sclerotic structure called the female sclerotized genital plate (FSGP) that is attached dorsally to the bursa, and of unknown use. The term ‘wings’ (W) describes the flat, fan-like projection extending to each lateral side of the FSGP, and the term ‘posterior bar’ refers to a curved sclerotized piece that sits posterior to the FSGP in some Eriauchenius. The male pedipalpal bulbs all have a conductor (C), and we believe what we are calling the conductor in this study is homologous to the conductor in Rix and Harvey (2011, 2012a, b), and to the bulb proapical process (BPAP) in Wood (2008). In some species there is an additional sclerite that we abbreviate SC that is likely an additional process of the conductor through a membranous connection. In some species we describe a median apophysis (MA) for convention, however we are not sure of the homology of this structure. We believe that the MA may be homologous to the ‘bulb dorsal sclerite’ (BDS) that was documented in Madagascarchaea gen. n. in Wood (2008). In Madagascarchaea gen. n. the sclerite that we call S1 was termed the ‘bulb lateral sclerite’ (BLS) in Wood (2008). The homology of archaeid pedipalpal parts is unclear and the terminology used here is not necessarily an argument for homology.

Taxonomy

ArchaeidaeKoch & Berendt, 1854

Diagnosis

Ecribellate, haplogyne, araneomorph spiders, three claws, with peg teeth, with cheliceral and pedicel stridulatory systems; modified carapace that wraps around the base of the chelicerae forming a constricted neck; extant genera with set of anterior booklungs and pair of posterior spiracles (configuration unknown in fossil archaeids). For complete description see Forster & Platnick (1984).

Discussion

While archaeids have a substantial fossil record (12 genera; Dunlop et al. 2017) from the Northern Hemisphere, the extant Southern Hemisphere genera form a well-supported monophyletic group based on morphology and molecular data (Wood et al. 2012; Wood et al. 2013). The extant clade can be distinguished from the fossil genera by the following combination of characters (Wood et al. 2012): presence of a tubercle on the sternum, which may be implicated in the pedicel stridulatory system that has been observed in the extant genera; the posterior edge of the carapace is truncated in the extant clades rather than tapering off as in the fossils; the distal portion of the chelicerae are curved towards the posterior; the presence of a brush of hairs on the prolateral side of the pedipalpal tarsi which interacts with cheliceral stridulatory file (Wood 2008), whereas fossils have only stridulatory picks (Madagascarchaeagracilicollis (Millot, 1948) is the exception, having both stridulatory picks and a brush of hairs). The extant archaeids are comprised of the following genera: Austrarchaea Forster & Platnick, 1984, and Zephyrarchaea Rix & Harvey, 2012a, from Australia; Eriauchenius, and Madagascarchaea gen. n., from Madagascar; and Afrarchaea Forster & Platnick, 1984, from South Africa. The Australian taxa (Austrarchaea and Zephyrarchaea) can be distinguished from the African and Madagascan taxa by the numerous spermathecae in females, the lack of a FSGP that sits immediately dorsal to the bursa, and the long wiry embolus on the male pedipalps (Forster and Platnick 1984; Rix and Harvey 2011, 2012a, b; Wood 2008). Zephyrarchaea can be distinguished from Austrarchaea by the carapace height to carapace length ratio being less than 2, and by the presence of accessory setae on or adjacent to the proximal cheliceral paturon bulge in males (Rix and Harvey 2012a). In females of the African and Madagascan genera there is a bursa with clusters of secretory pores, typically on the anterior and dorsal side of the bursa, and a FSGP is also present immediately adjacent to the dorsal side of the bursa.

GenusEriaucheniusO. P.-Cambridge, 1881

Type species

E.workmani O. P.-Cambridge, 1881, by original designation.

Diagnosis

Distinguished from the Australian genera by lacking spermathecae in the female genitalia, and instead having a bursa with secretory poreplates and a FSGP, and, in males, lacking the long wiry embolus on the male pedipalps seen in the Australian species. Distinguished from Afrarchaea by lacking the perpendicular keel on the FSGP, and from Madagascarchaea gen. n. by having 2–4 spines on the apex of the cephalon instead of 6, and lacking the retrolateral apophysis on the male pedipalpal patella.

Description

Total length 1.64–6.72. Carapace reddish-to-orangish brown with many white setae on small tubercules, organized in branching rows (see figs 5D, 6B in Wood 2008); pars cephalica elongated forming ‘head’ (distal portion of elongation) and ‘neck’ (constricted portion of elongation), with carapace tilt height divided by carapace length 1.44–2.85, with the angle of carapace tilt 54.7°–88.1°; with a pair of posterior, and often with an anterior pair, of pronounced-to-rudimentary protrusions on apex of cephalon or ‘head,’ each with a small-to-large spine; neck with fissure on anterior side running from chelicerae bases to labrum (see fig. 6B in Wood 2008). AME on a small-to-large bulge with a point or rounded at apex (see fig. 8C in Wood 2008). AME diameter larger than all other eyes; median ocular quadrangle (MOQ) wider in front than behind or than long; lateral eyes contiguous; sometimes with a short spine between median eyes and LE (see figs 8C and 18 in Wood 2008). Sternum reddish-to-orangish brown and longer than wide, hollowed out around coxae, with a border; setae with tuberculate bases; with expanded tubercle on posterior part of sternum, close to 4th coxae (see fig. 9F in Wood 2008), possibly implicated in the abdomen-petiole stridulatory system. Long sclerite between coxae and carapace (see fig. 7C in Wood 2008). Endites converging; serrula strongly pointed; labrum with two lateral projections on dorsal surface (see fig. 6A in Wood 2008). Small round chilum sclerite next to each cheliceral base. One triangular sclerite between and posteriad to the cheliceral bases, and an additional sclerite running along the length of the triangular sclerite base (see fig. 7A in Wood 2008). Chelicerae with a pronounced-to-rudimentary anterior protrusion with a downward or perpendicular pointing long-to-short spine, cheliceral spine/chelicerae length ratio 0.068–0.49; with stridulatory ridges on lateral side, and chelicerae curved to the posterior distal to the stridulatory ridges. The structure used in conjunction with the cheliceral stridulatory file appears to be a group of modified hairs on the prolateral side of the palp (Forster and Platnick 1984; Lotz 2003; Wood 2008); it is unknown whether sclerotized structures on the male palpal bulb are also used in conjunction with the cheliceral file, which has been observed in Madagascarchaea gen. n. (Wood 2008). Peg teeth in three rows; anterior row with two peg teeth and posterior row of two-three, both sitting opposite fang tip, median row of 18–30, strongest distally and gradually grading to normal setae. Teeth on retromargin 2–7, may have different numbers of teeth per chelicera on same individual.

Abdomen rounded in the “bourgini” (Figs 9–22) or triangular in the “workmani group” (Figs 3–8) due to a single dorsal protuberance, which can be small-to-large; containing numerous small, round, pale indentations throughout and with dark brown to purplish pigment throughout; covered in white-to-brown, thick setae; epigynum and booklung covers flat, sclerotized plates, which are not fused in females, but form a single fused epigastric ventral plate in males; abdominal dorsal plate with ridges (see fig. 5B in Wood 2008), sometimes fused with epigastric plate only in males; dorsal plate sometime extends along the anterior face of the abdomen (scutum) in males only; dorsal plate is always confined to the abdominal petiole region in females. Posterior respiratory system with two spiracular openings (see fig. 5C in Wood 2008).

Spinnerets surrounded by ring; rudimentary-to-fleshy colulus present. The following spinneret description is taken from examining published images of E.workmani (Griswold et al. 2005: figs 21–22): anterior lateral spinneret (ALS) spinning field divided, with one large and one smaller major ampullate gland (MAP) spigot on the median side and with approximately 42 smaller piriform gland (PI) (fewer in male) spigots on the lateral side. Posterior median spinneret (PMS) of female with one large median minor ampullate gland spigot (mAP), one lateral medium-sized aciniform gland (AC) spigot, and two lateral cylindrical gland (CY) spigots; the male PMS is the same except in lacking the CY. Posterior lateral spinnerets (PLS) with a middle row of five AC spigots; in females only, this row is flanked on the anterior side by two CY spigots and on the posterior side by one CY spigot.

Legs reddish or orangish to light brown, often with dark brown bands throughout, but especially on the tibia; covered sparsely with setae; ratio 1-2-4-3 or 1-4-2-3, typically 1-2-4-3 for species found in vegetation and 1-4-2-3 for species found in forest litter; one or two anterior rows of scopulae present on leg I, sometimes also present on leg II, and sometimes with a posterior row as well; metatarsus III and IV with a ventral cluster of modified hairs; femur IV distinctly curved (see fig. 7D in Wood 2008); femur I length 1.48–6.48 times the length of the carapace. Female palp with single claw.

Male pedipalpal femur, patella, tibia and cymbium without apophyses, however a cluster of spine like setae with enlarged bases occurs on the distal retrolateral side of the femur in some “bourgini group” species. Palpal bulb very diverse in shape, forming an enclosed pit that the conductor wraps around in the “workmani group” (Fig. 8D–L) and the “bourgini group – enclosed embolus group” (Fig. 9D–L), and with the embolus exposed and encircled by the conductor in the “bourgini group – exposed embolus group” (Fig. 13D–K). Embolus is heavily sclerotized, unconcealed, and wide-to-thin in the “bourgini group” and is sclerotized only at the tip and recessed in the “workmani group”. Conductor either a small dark ridged spiral that goes around the apex of the bulb and terminates in a triangular point (Fig. 9J–K), or a larger, wider piece that circles around the embolus, often with processes (Fig. 13D–I). In the “workmani group” the MA is present and is hidden behind the embolus in unexpanded bulbs (Fig. 8D–E, H); in most “bourgini group” species the male pedipalpal bulb has only a conductor and embolus, however, in some species a MA is also present (Figs 9E, H, 11D–I, 12D–F, 20D–I); in all “workmani group” species and in some “bourgini group” species there is an additional sclerite (SC) present on the male pedipalpal bulbs that may be an additional process on the conductor instead of a separate sclerite (Figs 8D–E, G–H, 9E–I).

Female genitalic bursa height in “workmani group” greater than bursa width (Fig. 8B), and in “bourgini group” less than or equal to bursa width (Fig. 11B); bursa with secretory poreplates with pores distributed in a small continuous group (Fig. 17C) or in a large discontinuous clumpy group on each side (Fig. 8B), although poreplates are absent in E.bourgini (Millot, 1948) and E.zirafy sp. n. that instead have a sclerotized invagination on either side of the bursa (Fig. 13C); with a dorsal sclerotized plate (FSGP) that can be either a simple arched piece that is wider than long (Fig. 15C) or a more elaborate piece that can have a posterior extension, or points at the top, and that can be longer than wide (Fig. 14B); FSGP in most species with wing-like projections extending to each lateral side that can be reduced-to-large, and that can be heavily sclerotized-to-translucent (Fig. 8B). All species of the “bourgini group” have a large-to-small curved piece that sits posterior to the epigastric furrow, termed the “posterior bar” (Figs 13B, 16B), not present in the “workmani group”. Legendre (1967) suggested that the male palp might come into contact with the FSGP during copulation and that the FSGP may offer tactile information to the male or female. Alternatively, the FSGP may serve as an anchor for muscle attachment (Griswold et al. 2005).

Distribution

Madagascar.

Discussion

Eriauchenius contains two main clades, the “workmani group” and the “bourgini group” (Fig. 1). The “workmani group” is distinctive: the abdomen has a single tubercle making it triangular in shape (Fig. 8A), whereas the “bourgini group” has a rounded abdomen (Fig. 9A); the “workmani group” also has a highly elongated, constricted “neck” (Fig. 8A). Furthermore, in the “workmani group” the bursa height is greater than the bursa width (Fig. 8B), whereas in the “bourgini group” the bursa height is less than or equal to the bursa width and there is a posterior bar on the FSGP (Fig. 11B–C). The “workmani group” species are bigger in general than other Madagascan archaeid species, ranging in body length from 3.24–6.72.

The “bourgini group” is further broken up into two groups, the “enclosed embolus group” and the “exposed embolus group.” Unfortunately, specimens from the “enclosed embolus group” were not included in the phylogeny of Wood et al (2015), so it is currently unknown whether this group is monophyletic and what their phylogenetic relationship is to other archaeids. The “embolus enclosed group” contains four species (E.fisheri, E.goodmani sp. n., E.harveyi sp. n., E.sama sp. n.) that share the following traits: the male pedipalpal bulb is enclosed (Figs 9D–L, 10D–F, 11D–K, 12D–F), the female bursa is sclerotized and covered in pores on the ventral side (Figs 9C, 10C, 11C, 12C) as opposed to the anterior or dorsal side in other Eriauchenius, and the AME are virtually flush with the cuticle (Fig. 11L), rather than being on bulges. In the “exposed embolus group” the male pedipalpal bulbs have a more open form, with the embolus exposed and encircled by the elongated conductor, and there is also a membraneous sac on the bulb that is adjacent to the embolus base (Fig. 17D–L). Typical “bourgini group” pedipalpal bulbs have only an embolus and a conductor, and no other sclerites, the exceptions are E.fisheri, E.harveyi sp. n., E.rixi sp. n., and E.wunderlichi sp. n., which also have a MA and SC, and E.goodmani sp. n., which also has a SC.

E.fisheri and E.mahariraensis were transferred to Eriauchenius from Afrarchaea because they lack the FSGP keel observed in Afrarchaea and because the FSGP has a posterior bar. E.mahariraensis was included in the phylogenetic analysis of Wood et al (2015) and fell inside Eriauchenius with a branch support posterior probability value of 1.0. Although, E.fisheri was not including in this phylogenetic analysis, the morphological evidence supports this transfer.

Key for Eriauchenius

1

In males and females, abdomen triangular, with a single tubercule on the dorsal side (Fig. 8A); in females, genitalia with bursa height greater than bursa length, posterior bar absent (Fig. 8B)

2, “workmani group”

–

Abdomen rounded in both sexes, without a tubercle (Fig. 11A); in females, bursa height less than or equal to bursa width, posterior bar present (Fig. 11B–C)

Male pedipalpal bulbs with apical conductor encircling a pit-like cavity (Figs 9D–K, 12D–F); conductor terminates in two pieces on the retrolateral side, one piece rounded and the other piece tapering off to a point (except blunt in E.goodmani sp. n.) (Figs 9D–I, 12F); in females, the ventral side of the bursa is sclerotized and covered in pores (Figs 9C, 10C, 11C, 12C) as opposed to the anterior side

12, “enclosed embolus group”

–

Male pedipalpal bulbs with a large embolus, encircled by conductor, with a white membraneous sac of cuticle that sits close to the base of the embolus (Figs 13D–J, 17D–L, 19D–L, 20D–L); conductor is large and may have projections (Figs 13D–K, 19D–I); in females, the secretory pores are in two groups on the anterior side of the bursa (Figs 17C, 19C) or there are two sclerotized invaginations on the anterior side of the bursa (Figs 13C, 22B–C)

15, “exposed embolus group”

“Enclosed embolus group”

12

In males, embolus is long and narrow, wire-like (Figs 9E, G–H, L, 11D–I); in females, “wings” on FSGP are reduced (Figs 9B, 11B)

In males, embolus tip with two curves making an “s” shape (Fig. 9E, H, K–L)

E.fisheri

14

In male pedipalpal bulb, the apical portion of the tegulum, where the conductor swirls around, is elongated so that the bulb is almost twice as long as it is wide (Fig. 12D, F); SC sclerite as wide and nearly as long as embolus (Fig. 12D–E); in females, posterior bar on genitalia has a large bulge in the center (Fig. 12B)

E.wunderlichi

–

In males, apical portion of the tegulum not as elongated (Fig. 10D, F); embolus wider and longer than SC sclerite (Fig. 10D–E); in females, posterior bar on genitalia lacking large bulge in the center (Fig. 10B)

E.goodmani

“Exposed embolus group”

15

In both males and females, posterior pair of spines on the apex of the cephalon on large protrusions (Figs 13A, 22A)

In males, pedipalpal bulb shape with the embolus and conductor originating from the posterior portion of the bulb (Fig. 19D–L); in females, FSGP with two lateral bulges (Fig. 19B)

E.ratsirarsoni

–

Male pedipalpal bulb shape with the embolus and conductor originating from the center of the bulb (Fig. 21D–L); FSGP lacking two lateral bulges (Fig. 21B)

E.sama

22

Male pedipalpal bulb posterior half-moon shaped, although there is variation in degree (compare tegulum shape in Fig. 17J–L with Fig. 17D–I), and constriction on conductor with a ridge (Fig. 17H); in females, posterior bar reaches the anterior edge of the FSGP (Fig. 17B)

E.milloti

–

Posterior edge of male pedipalpal bulb more rounded (Fig. 14D–E, G–I) and lacking a ridge on the conductor constriction (Fig. 14H); in females, posterior bar less than half of the length of the FSGP (Fig. 14B)

E.lukemacaulayi

–

Males unknown; in females, FSGP with a broad, rectangular posterior elongation, and with posterior bar more than half, but less than the full length of the FSGP (Fig. 16B)

Etymology

The specific name is a noun in apposition and commemorates King Andriamanelo, the founder of the Merina Kingdom.

Diagnosis

Males and females are considered part of the “workmani group” based on having a single dorsal protuberance on the abdomen (a triangular shaped abdomen). Males are distinguished from all other species in the “workmani group” by the tip of the conductor, which is divided into two separate sclerotized processes (Fig. 3E, H, K), and by the hook shaped MA (Fig. 3C). Females are distinguished from all other species in the “workmani group” by the presence of a heavily sclerotized “T” shaped structure on the posterior of the bursa (Fig. 3B, L).

Natural history

Specimen were collected in montane rainforest, rainforest and tropical dry forest, by beating low vegetation, by beating and sweeping forest understory, and by general collecting day and night. Specimens occur from 30–1300 m in elevation.

Etymology

The specific name is a noun in apposition and commemorates King Andrianampoinimerina, who unified the Merina Kingdom.

Diagnosis

Males and females are considered part of the “workmani group" based on having a single dorsal protuberance on the abdomen (a triangular shaped abdomen) (Fig. 4A). Males and females are distinguished from all other species in the “workmani group" by the strong abdomen markings and by the “neck” being very upright, having a tilt angle that is greater than 80° (Fig. 4A). Males are distinguished from all other “workmani group” species except E.rafohy sp. n. by having the MA with a bifurcation (Fig. 4C).

Description

Male holotype (CASENT9018910, from Montagne d’Anjanaharibe, Madagascar). Total length 4.32, carapace 1.52 long, 1.29 wide. Abdomen 2.40 long, 1.73 high, with a dorsal hump. Carapace tilt angle 88.1°, tilt height (CtH) 3.94, constriction 0.35, head length 1.32, neck length 2.34. CtH divided by carapace length 2.59. Cephalon with AME on a large bulge and 4 post-ocular protrusions on the apex of the cephalon (Fig. 4A), each provided with a short modified spine at the tip. Chelicerae 3.71 long, and with spine 0.33 from base of chelicerae (Fig. 4A). Femur I 9.58 long. Sternum 1.08 long, 0.50 wide. Carapace, chelicerae, sternum and femora I & II reddish dark brown with many white setae. Legs I & II with lighter annulations close to leg joints. All coxae yellowish brown and legs III & IV yellowish brown, with darker annulations on femora, tibiae and metatarsi. Abdomen with characteristic color pattern of undulating brown ‘rings’, with tufts of white setae and white book-lung covers (Fig. 4A). Pedipalpal tegulum of the “workmani group” form, with apical conductor encircling a pit-like cavity (Fig. 4D–I). Conductor tip is a broad triangular point (Fig. 4D–I) similar to other “workmani group” species except E.andriamanelo. MA with a bifurcation (Fig. 4C). Embolus similar to other “workmani group” species, being broad and complex with the sperm duct opening in the middle and sclerotization only at the tip (Fig. 4D–E, G–H).

Natural history

Specimens were collected in montane rainforest, rainforest, and montane shrubland, by beating low vegetation, beating vegetation, sweeping, raking, and by general collecting day and night. Specimens were collected from 195–2000 m in elevation.

Etymology

The specific name is a noun in apposition and commemorates Queen Rafohy.

Diagnosis

Males and females are considered part of the “workmani group” based on having a single dorsal protuberance on the abdomen (a triangular shaped abdomen) (Fig. 5A). Males are distinguished from the “workmani group” species E.andriamanelo sp. n., E.ranavalona sp. n., and E.rangita sp. n. by having a bifurcating MA (Fig. 5C), from E.andrianampoinimerina by lacking the distinctive abdomen markings (Fig. 5A), and E.workmani by lacking the large bump on the pedipalpal bulbs (Fig. 5D–K). Females are distinguished from E.andriamanelo by lacking the heavily sclerotized “T” shaped structure on the posterior of the bursa, from E.andrianampoinimerina by lacking the strong abdomen markings and by the “neck” having a tilt angle that is less than 80°, and from E.ranavalona by lacking the lime-green abdomen markings. Females are indistinguishable from the remaining “workmani group” species.

Natural history

Specimens have been collected in montane rainforest through general collecting, beating vegetation, sweeping, raking, cryptic searching, and among fallen logs and litter in altitudes from 1300–1638 m above sea level. One specimen was collected with a hatched eggsac.

Distribution

Known only from Antananarivo Province in central Madagascar (Fig. 30).

Etymology

The specific name is a noun in apposition and commemorates Queen Ranavalona III, the last sovereign of the Kingdom of Madagascar before it became a French colony.

Diagnosis

Males and females are considered part of the “workmani group” based on having a single dorsal protuberance on the abdomen (a triangular shaped abdomen) (Fig. 6A). Males and females are distinguished from all other species in the “workmani group” by the lime-green abdomen markings in living specimens (Fig. 6C–D), which is sometimes faded in alcohol preserved specimens, and in males, by the non-bifurcating MA that is broad and tapers to a point (Fig. 6H–I), rather than being hook shaped as seen in E.andriamanelo (Fig. 3C).

Description

Male holotype (CALENT9010047, from Parc Nationale Ranomafana, Madagascar). Total length 3.53, carapace 1.48 long, 1.28 wide. Abdomen 1.93 long, 2.23 high, with a dorsal hump. Carapace tilt angle 75.1°, tilt height (CtH) 3.25, constriction 0.44, head length 1.03, neck length 1.83 (Fig. 2). CtH divided by carapace length 2.20. Cephalon with AME on a large bulge and 4 post-ocular protrusions on the apex of the cephalon (Fig. 6A), each provided with a short modified spine at the tip. Chelicerae 3.28 long, and with spine 0.42 from base of chelicerae (Fig. 6A). Femur I 5.76 long. Sternum 0.92 long, 0.55 wide. Carapace, chelicerae, sternum and femora I & II reddish dark brown with many white setae, and lighter brown areas on head, neck and chelicerae. All coxae yellowish brown and legs III & IV yellowish brown, with darker annulations on femora, tibiae and metatarsi. Abdomen mostly mottled brown with tufts of white setae, white book-lungs, and characteristic lime-green posterior-dorsal area in living species (Fig. 6C–D; often faded to yellowish-white in ethanol preserved material). Pedipalpal tegulum of the “workmani group” form, with apical conductor encircling a pit-like cavity (Fig. 6E–J). Conductor tip is a broad triangular point similar to other “workmani group” species except E.andriamanelo (Fig. 3K), where the conductor is divided. MA without a bifurcation, broad, and tapering toward the tip (Fig. 6E–F, H–I). Embolus similar to other “workmani group” species, being broad and complex with the sperm duct opening in the middle and sclerotization only at the tip.

Natural history

Specimens have been collected in montane rainforest and evergreen secondary rainforest through beating vegetation, including clumps of dead, dry foliage, by beating low vegetation, and general collecting day and night. Specimens were collected from 900–1200 m above sea level.

Distribution

Known only from Ranomafana National Park in southwestern Madagascar (Fig. 30).

Etymology

The specific name is a noun in apposition and commemorates Queen Rangita.

Diagnosis

Males and females are considered part of the “workmani group” based on having a single dorsal protuberance on the abdomen (a triangular shaped abdomen) (Fig. 7A). Males are distinguished from all other species in the “workmani group” by the shape of the MA that has two bifurcations, one deep and one shallow (Fig. 7C). Females are distinguished from E.andriamanelo by lacking the heavily sclerotized “T” shaped structure on the posterior of the bursa, from E.andrianampoinimerina by lacking the strong abdomen markings and by the “neck” having a tilt angle that is less than 80°, and from E.ranavalona by lacking the lime-green abdomen markings. Females are indistinguishable from the remaining “workmani group” species.

Description

Male holotype (CASENT9010053, from Ambohimanga Village, Madagascar). Total length 4.15, carapace 1.58 long, 1.37 wide. Abdomen 2.44 long, 2.74 high, with a prominent dorsal hump. Carapace tilt angle 71.52°, tilt height (CtH) 3.64, constriction 0.50, head length 1.50, neck length 2.09. CtH divided by carapace length 2.30. Cephalon with AME on a large bulge and 4 post-ocular protrusions on the apex of the cephalon (Fig. 7A), each provided with a short modified spine at the tip. Chelicerae 3.69 long, with seta 0.52 from base of chelicerae (Fig. 7A). Femur I 6.93 long. Sternum 1.11 long, 0.59 wide. Carapace, chelicerae, sternum, coxae and femora I & II reddish dark brown with many white setae. Legs III & IV yellowish brown, with darker annulations on femora, tibiae and metatarsi. Abdomen yellowish brown, mottled with grayish brown, and light brown book-lung covers, all covered with many white setae (Fig. 7A). Pedipalpal tegulum of the “workmani group” form, with apical conductor encircling a pit-like cavity (Fig. 7D–K). Conductor tip is a broad triangular point similar to other “workmani group” species except E.andriamanelo (Fig. 3H), where the conductor is divided. MA with a broad unsclerotized transparent base and two apical sclerotized processes, one forked and the other un-forked, so the MA looks tri-forked in the unexpanded palp (Fig. 7C, G, J–K). Embolus similar to other “workmani group” species, being broad and complex with the sperm duct opening in the middle and sclerotization only at the tip (Fig. 7E, H, K).

Natural history

Specimens have been collected in montane rainforest through general collecting day and night, ‘cryptic searching’, and within a fallen palm frond, at altitudes of 1050–1550 m above sea level. Several specimens have been collected with separate eggsacs that are almost the size of the spider and contains 30–40 relatively large white eggsacs wrapped in thin transparent silk. A couple of hatched eggsacs have also been collected.

Taxonomical note

Originally named Eriaucheniusworkmanni by Pickard-Cambridge in 1881, but subsequently referred to as Eriaucheniusworkmani, except for a very recent reference to the original spelling (World Spider Catalogue 2017). Article 33.3.1 of the International Code of Zoological Nomenclature states that “when an incorrect subsequent spelling is in prevailing usage and is attributed to the publication of the original spelling, the subsequent spelling and attribution are to be preserved and the spelling is deemed to be a correct original spelling”. We here follow the World Spider Catalogue and consider the spelling “workmani” as “prevailing use” and therefore the correct spelling of this species name.

Diagnosis

Males and females are considered part of the “workmani group” based on having a single dorsal protuberance on the abdomen (a triangular shaped abdomen). Males of E.workmani are distinguished from all other species by the large bump on the pedipalpal bulbs (see arrow in Fig. 8H). Females are distinguished from E.andriamanelo by lacking the heavily sclerotized “T” shaped structure on the posterior of the bursa, from E.andrianampoinimerina by lacking the strong abdomen markings and by the “neck” having a tilt angle that is less than 80°, and from E.ranavalona by lacking the lime-green abdomen markings. Females are indistinguishable from the remaining “workmani group” species.

Natural history

Specimens have been collected in montane and lowland rainforest through general collecting, beating vegetation, sweeping, raking, cryptic searching, and among fallen logs and litter at altitudes of 26–1200 m above sea level. Several specimens were collected with eggcases that look similar to eggcases from other “workmani group” species.

Distribution

Widely distributed in the eastern rainforests of Madagascar from north to south (Fig. 29).

Nomenclatural remarks

The holotype is a juvenile. We assume it to be the species we are describing because this species is the most widespread common species of the “workmani group”. Millot (1948) assumed the same thing.

“Workmani group” spp. – unidentified specimens

There were many "workmani group" specimens that could not be confidently identified to a known species, listed below, due to the absence of adult male specimens and molecular data. Additionally, there was one penultimale male specimen that died while molting in captivity that was included in the molecular phylogeny of Wood et al. (2015). This specimen is likely a new species based on the phylogenetic relationships (shown as Eriauchenius sp. 1 in Fig. 1). However, since there are no known adult males, and only one adult female specimen, we chose not to describe this species at this time: 1 penultimate male, MADAGASCAR, Fianarantsoa, Parc National Andrigitra, 34 km S Ambalavao, 22°09'24.9"S, 46°57'08.7"E, 1830 m, 8–9 Jan 2009, primary montane rainforest, general collecting day, beating vegetation, H. Wood (USNMENTO01377166); 1F, Madagascar, Fianarantsoa, Parc National Andrigitra, 34 km S Ambalavao, 22°09'24.9"S, 46°57'08.7"E, 1830 m., 8–9 Jan 2009, primary montane rainforest, general collecting day, beating vegetation, H. Wood (USNMENT01377165).

Diagnosis

Males are distinguished from other “bourgini group” species except E.harveyi sp. n., E.goodmani sp. n., and E.wunderlichi sp. n., by having the pedipalpal tegulum of the “workmani group” form, with the apical conductor encircling a pit-like cavity (Fig. 9D–L). Males are distinguished from E.goodmani sp. n., E.harveyi sp. n. and E.wunderlichi sp. n. by having an s-shaped embolus (Fig. 9E, H, K–L). Females are distinguished from the “bourgini group” except for E.goodmani sp. n. and E.wunderlichi sp. n. by having a bursa with two large groups of poreplates on a sclerotized plate that covers the ventral side of the bursa (Fig. 9C). In contrast, in other “bourgini group” species the poreplates are in smaller clusters on the anterior side of the bursa. Females are distinguished from E.goodmani sp. n. by having a thinner posterior bar (Fig. 11C), and from E.wunderlichi sp. n. by lacking the large bulge in the center of the posterior bar (Fig. 12B). Females are indistinguishable from E.harveyi sp. n.

Description

Female holotype (CASENT9012340, from Reserve Andringitra, Madagascar). This specimen was damaged during shipping so that the “head” is broken off from the “neck,” rendering some measurements impossible. Total length 2.38, carapace 1.11 long, 0.98 wide. Abdomen 1.16 long, 1.17 high. Carapace tilt height (CtH) 1.61, head length 0.98, carapace tilt angle, carapace constriction, and neck length unknown due to damage. CtH divided by carapace length 1.45. Cephalon with AME virtually flush with surrounding cuticle, and with a single pair of short modified spines at the apex (Fig. 9A). Chelicerae 1.86 long, and with a long spine 0.21 from base of chelicerae and projecting perpendicular to the cheliceral cuticle. Femur I 1.73 long. Sternum 0.72 long, 0.47 wide. Carapace, chelicerae, sternum and legs reddish brown with white setae, but reduced numbers compared to other Eriauchenius. All patella lighter in color, being more yellowish white. Abdomen mottled brown and beige, with tufts of white setae, although reduced in number compared to other Eriauchenius (Fig. 9A). Genitalia with a noncomplex FSGP, with posterior bar (not visible in Fig. 9B, but similar to Fig. 11B–C), and with “wings” reduced and nearly transparent, with poreplates in two large groups, divided down the middle and on a sclerotized plate that covers the ventral side of the bursa (Fig. 9C, similar to Fig. 11C).

Male paratype (CASENT9018939). Total length 2.18, carapace 1.01 long, 0.92 wide. Abdomen 1.13 long, 1.16 high. Carapace tilt angle 79.9°, tilt height (CtH) 1.53, constriction 0.65, head length 0.91, neck length 0.72 . CtH divided by carapace length 1.51. Cephalon as in female. Chelicerae 1.58 long, and with long spine 0.20 from base of chelicerae and projecting perpendicular. Femur I 1.64 long. Sternum 0.65 long, 0.44 wide. Colors as in female. Pedipalpal tegulum of the “workmani group” form, with apical conductor encircling a pit-like cavity (Fig. 9D–L). Conductor tip tapering off into a sharp point (Fig. 9F,I,L). MA trifurcating into three prongs (Fig. 9E, H, L), and with a sclerite (SC) that may be part of the conductor, thicker and more sclerotized than the transluscent thin structure seen in the “workmani group” (Fig. 9E, H). Embolus dark, wire-like and with two curves making an “s” shape (Fig. 9E, H, K–L).

Natural history

Specimens were collected in rainforest in the leaf litter from 1200–1990 m in elevation.

Distribution

Known only from around Andringitra Massif in southeast Madagascar (Fig. 31).

Nomenclatural remarks

Distributions of E.fisheri and E.harveyi sp. n. are in close proximity, however, E.fisheri seems to occur at higher elevations than E.harveyi sp. n. Furthermore, E.fisheri is slightly larger than E.harveyi sp. n. Male and female conspecifics were associated based on these factors, but due to the small number of specimens available, this association may be incorrect. Future molecular work as well as additional collection of specimens from more localities can help resolve this issue.

Other material examined

Etymology

The specific name is a patronym to honor Dr. Steven Goodman, who collected the specimens and for his extensive work on Madagascar’s biodiversity.

Diagnosis

Males are distinguished from other “bourgini-group” species except E.harveyi sp. n., E.fisheri, and E.wunderlichi sp. n., by having the pedipalpal tegulum of the “workmani group” form (Fig. 10E), with the apical conductor encircling a pit-like cavity (Fig. 10D–E). Males are distinguished from E.harveyi sp. n., E.fisheri, and E.wunderlichi sp. n., by the conductor being greatly elongated and blunt at the tip (Fig. 10F). Females are distinguished from “bourgini group” species, except E.harveyi sp. n., E.fisheri, and E.wunderlichi sp. n., by having a bursa with two large groups of poreplates on a sclerotized plate that covers the ventral side of the bursa (Fig. 10C), whereas in other “bourgini group” species the poreplates are in smaller cluster on the anterior side of the bursa. Females are distinguished from E.harveyi sp. n. and E.fisheri, by having a thicker posterior bar on the internal genitalia (Fig. 10B), and distinguished from E.wunderlichi sp. n. females by lacking the large bulge in the center of the posterior bar (Fig. 12B).

Other material examined

Etymology

The specific name is a patronym to honor Dr. Mark Harvey for his work on the Australian archaeids.

Diagnosis

Males are distinguished from other “bourgini group” species except E.fisheri, E.goodmani sp. n. and E.wunderlichi sp. n. by having the pedipalpal tegulum of the “workmani group” form, with the apical conductor encircling a pit-like cavity (Fig. 11D–K). Males are distinguished from E.fisheri by having a comparably straighter embolus, with only a broad curve (Fig. 11E, H, J–K), rather than s-shaped, from E.goodmani sp. n. by lacking the elongated conductor that is broad and blunt at the tip (Fig. 10F), rather than tapering off to a fine point (Fig. 11F, I), and from E.wunderlichi sp. n. by lacking an elongation in the apical portion of the tegulum where the conductor swirls around making the bulb almost twice as long as it is wide (Fig. 12F). Females are distinguished from the “bourgini group” except for E.goodmani sp. n. and E.wunderlichi sp. n. by having two large groups of poreplates on a sclerotized plate that covers the ventral side of the bursa (Fig. 11C), whereas in other “bourgini group” species the poreplates are in smaller clusters on the anterior side of the bursa. Females are distinguished from E.goodmani sp. n. by having a thinner posterior bar (Fig. 11B), and from E.wunderlichi sp. n. by lacking the large bulge in the center of the posterior bar (Fig. 12B). Females are indistinguishable from E.fisheri.

Other material examined

Etymology

The specific name is a patronym to honor Jörg Wunderlich, for his work documenting fossil archaeids.

Diagnosis

Males are distinguished from other “bourgini group” species except E.harveyi sp. n., E.fisheri, and E.goodmani sp. n., by having the pedipalpal tegulum of the “workmani group” form, with the apical conductor encircling a pit-like cavity (Fig. 12D–F). Males are distinguished from E.harveyi sp. n., E.fisheri, and E.goodmani sp. n., by having an elongated apical portion of the tegulum, so that the bulb is almost twice as long as it is wide (Fig. 12F). Females are distinguished from the “bourgini group,” except from E.harveyi sp. n., E.fisheri, and E.goodmani sp. n., by having a bursa with two large groups of poreplates on a sclerotized plate that covers the ventral side of the bursa (Fig. 12B–C), whereas in other “bourgini group” species the poreplates are in smaller clusters on the anterior side of the bursa. Females are further distinguished from the “bourgini group” by having a large bulge on the center of the posterior bar (Fig. 12B, arrow).

Description

Male holotype (CASENT9062702, from Parc National Befotaka-Midongy, Madagascar). The abdomen of this specimen is missing making some measurements impossible. Carapace 1.11 long, 0.89 wide. Carapace tilt angle 76.7°, tilt height (CtH) 1.80, constriction 0.76, head length 1.02, neck length 0.87. CtH divided by carapace length 1.62. Cephalon with AME virtually flush with surrounding cuticle, and with 2 short modified spines at the apex. Chelicerae 1.78 long, and with long spine 0.19 from base of chelicerae and projecting perpendicular to the cheliceral cuticle (Fig. 12A). Femur I 1.98 long. Sternum 0.71 long, 0.42 wide. Carapace, chelicerae, sternum and legs reddish brown with white setae. Legs with darker annulations on tibiae and metatarsi. Abdomen color unknown. Pedipalpal tegulum of the “workmani group” form, with apical conductor encircling a pit-like cavity, however the apical portion of the tegulum is elongated so that the bulb is almost twice as long and wide (Fig. 12D–F). Conductor tip tapering off into a sharp point (Fig. 12F). MA rounded and small, and nearly hidden behind sclerite SC and embolus. Sclerite SC long and heavily sclerotized and in close proximity and running parallel to the embolus. Embolus dark, thick, and with a swirl at the base and blunt at the tip (Fig. 12E–D).

Variation

Natural history

Specimens were collected in montane rainforest from 1250–1270 m in elevation by sifting litter.

Distribution

Known only from southeast Madagascar (Fig. 31).

Nomenclatural remarks

The male and female of E.wunderlichi sp. n. occur in rainforest areas in southwestern Madagascar that are not contiguous. The male and female may not be conspecifics, however based on morphology they likely belong in the “enclosed embolus group.” Future molecular work as well as additional collection of specimens from more localities can help resolve these issues.

Diagnosis

Males and females are distinguished from other Eriauchenius by having pointed extensions on coxae I (Fig. 13L, arrows). Males and females of E.bourgini and E.zirafy sp. n. are also distinguished by having two large protrusions on the crown of the cephalon (Fig. 13A). Females of E.bourgini and E.zirafy sp. n. are indistinguishable but are different from all other Eriauchenius by having two sclerotized invaginations on the bursa (Figs 13B, 22B, arrows). In E.bourgini males the conductor has four elaborate processes (Fig. 13D–K), and in and E.zirafy sp. n. males, five elaborate processes (Fig. 22D–K).

Description

Male (based on CASENT9001207, from Réserve Spéciale d’Ambohitantely, Madagascar). Total length 1.94, carapace 0.79 long, 0.74 wide. Abdomen 1.09 long, 0.97 high. Carapace tilt angle 72.1°, tilt height (CtH) 1.80, constriction 0.33, head length 0.89, neck length 0.93. CtH divided by carapace length 2.28. Cephalon with AME on a large bulge and 4 post-ocular spines on the crown of the cephalon, with the posterior pair on large protrusions and the anterior pair not on protrusions, and with 1 small spine between the LE and median eyes (on each side, for a total of 2). Chelicerae 1.68 long, and with a short spine 0.70 from base of chelicerae, projecting downward. Femur I 2.13 long. Sternum 0.51 long, 0.36 wide. Carapace, chelicerae, and sternum dark reddish brown with white setae. Coxae and legs lighter brown, with darker annulations on tibiae and metatarsi. Coxae I with posterior extensions (Fig. 13L). Abdomen mottled brown and beige, with a bright white patch on each lateral side, with tufts of white setae (as in Fig. 22A from E.zirafy sp. n.). Pedipalpal bulb with a small membraneous sac on the retrolateral side, with the base of the conductor small and triangular (labeled “c” in Fig. 13D, F–G), and with the remainder of the conductor wrapping around the embolus and with four long processes (Fig. 13D–F). The embolus is thick and contains several processes.

Etymology

The specific name is a patronym to honor Dr. Luke Macaulay for his help collecting palpimanoid spiders.

Diagnosis

Male is distinguished from other “bourgini group” species except E.milloti sp. n. and E.pauliani (and presumably E.milajaneae sp. n.) by the presence of a lateral process on the conductor (Fig. 14G–I, asterisk), which is narrow in E.lukemacaulayi and E.milloti (Fig. 14H). Males are distinguished from E.milloti sp. n. by lacking the quarter-moon-like-shape of the palpal bulb in E.milloti (Fig. 17D–L). Eriaucheniuslukemacaulayi sp. n., E.milajaneae sp. n., and E.milloti sp. n. (and presumably E.pauliani) females are distinguished from other “bourgini group” species by the enlargement of the posterior bar. While this bar is present in other “bourgini group” species, in E.lukemacaulayi sp. n., E.milajaneae sp. n., and E.milloti sp. n. this bar is large and curved into a u-shape. In E.lukemacaulayi sp. n., either end of this bar extends anterior just past the bottom of the “wings” (Fig. 14B), whereas in E.milajaneae sp. n. and E.milloti sp. n. the curved bar goes at least to the middle of the “wings” (Figs 16B, 17B). Eriaucheniuslukemacaulayi sp. n. females are also distinguished by the rounded posterior elongation of the FSGP that curves to the dorsal (Fig. 14F, arrow).

Description

Male holotype (USNMENT01377191, from Parc National Andrigitra, Madagascar). Total length 2.38, carapace 1.05 long, 0.97 wide. Abdomen 1.30 long, 1.47 high. Carapace tilt angle 63.59°, tilt height (CtH) 2.29, constriction 0.55, head length 1.03, neck length 1.04. CtH divided by carapace length 2.18. Cephalon with AME on a large bulge and 4 post-ocular spines on the crown of the cephalon, not on protrusions, and with 1 small spine between the LE and median eyes (on each side, for a total of 2). Chelicerae 1.99 long, and with a spine 0.27 from base of chelicerae and projecting perpendicular to the cheliceral cuticle. Femur I 2.56 long. Sternum 0.66 long, 0.44 wide. Carapace, chelicerae, sternum, and legs reddish brown with white setae. All legs with parts that are lighter brown and parts with darker annulations. Abdomen dark brown mottled with yellowish white spots and interspersed with white setae. Pedipalpal bulb with a membranous sac above the base of the embolus on the retrolateral side, with a greatly exposed embolus that is encircled by the conductor as conductor tapers off (Fig. 14D–I). Conductor with a constriction, followed by a narrow, curved process (Fig. 14G–I, asterisk) and a bulge (Fig. 14H, arrow). Embolus broad and dark (Fig. 14G–I).

Female paratype (USNMENT01377192). Total length 2.42, carapace 1.00 long, 0.93 wide. Abdomen 1.34 long, 1.56 high. Carapace tilt angle 65.6°, tilt height (CtH) 2.18, constriction 0.53, head length 0.97, neck length 1.04. CtH divided by carapace length 2.18. Cephalon as in male. Chelicerae 2.04 long, and with short spine 0.37 from base of chelicerae and projecting downward (Fig. 14A). Femur I 2.40 long. Sternum 0.63 long, 0.44 wide. Colours as in male, expect anterior of abdomen dark brown with yellowish spots and posterior of abdomen mostly yellowish white with some white patches (Fig. 14A). Female genitalia FSGP with a posterior elongation that curves dorsally (Fig. 14F, arrow), and with lateral bulges at the base of each “wing” (Fig. 14B), with a large U-shaped posterior bar that extends just past the posterior portion of the “wings” (Fig. 14B), with poreplates in one group on each lateral side of the bursa anterior (Fig. 14C).

Diagnosis

Males and females are distinguished from other Eriauchenius, except E.ratsirarsoni and E.sama sp. n., by the presence of 4 small spines on the apex of the cephalon, and having the cheliceral spine pointing perpendicular (as in Fig. 11L). In the holotype female the cheliceral spine is broken off, but the socket shape suggests the spine would be at a perpendicular orientation as it is in the paratype male. Eriaucheniusmahariraensis is also distinguished from other Eriauchenius males, by having membranous tissue close to the conductor tip (Fig. 15E–L), and other Eriauchenius females by having a very reduced FSGP with the width divided by height greater than 2 (Fig. 15B).

Description

Female holotype (MCZ, from Parc National Ranomafana, Madagascar). Total length 1.91, carapace 0.74 long, 0.69 wide. Abdomen 1.06 long, 1.24 high. Carapace tilt angle 64.4°, tilt height (CtH) 1.32, constriction 0.44, head length 0.68, neck length 0.64. CtH divided by carapace length 1.77. Cephalon with AME on a small bulge. Cephalon with 4 small post-ocular spines (one is missing or broken off) on the crown of the cephalon, not on protrusions, and 1 spine between the LE and AME (on each lateral side, for a total of 2, but broken off on the right side). Chelicerae 1.33 long, and with a spine 0.24 from base of chelicerae, however, the spines are broken off and only the socket remains; the socket shape suggests the spines would projecting perpendicular to the cheliceral cuticle. Femur I 1.46 long. Sternum 0.51 long, 0.35 wide. Carapace, chelicerae, sternum, and legs reddish brown with white setae; patellas lighter brown. Abdomen anterior dark brown with light circular patches and posterior light tan; abdomen with white and brown setae (Fig. 15A). Female genitalia with a small and simple FSGP, with “wings” highly reduced; posterior bar is present, but the dissection was such that it remains attached to the abdomen and is not visible in Fig. 15B–C; with poreplates in one small group on each lateral side of the bursa anterior, and a small sclerotized piece on the ventral side of the poreplates (Fig. 15C, arrow).

Male paratype (USNMENT01377173). Total length 1.64, carapace 0.78 long, 0.68 wide. Abdomen 0.83 long, 0.97 high. Carapace tilt angle 66.7°, tilt height (CtH) 1.31, constriction 0.44, head length 0.72, neck length 0.61. CtH divided by carapace length 1.69. Cephalon with AME on a small bulge, and with 4 small post-ocular spines (one is missing or broken off) on the crown of the cephalon, and 1 spine between the LE and AME (on each side, for a total of 2). Chelicerae 1.23 long, and with small spine 0.17 from base of chelicerae (Fig. 15A). Femur I 1.63 long. Sternum 0.49 long, 0.32 wide. Cephalothorax colors as in female. Abdomen mostly light tan, but mottled with dark brown areas that have light circular patches; with white and brown setae; posterior of abdomen with bright white areas. Pedipalpal bulb with a small membraneous sac above the embolus base, with a greatly exposed broad and dark embolus that is encircled by the conductor (Fig. 15D–L). Conductor with a membranous area at the distal end that swirls around the embolus (Fig. 15D–L).

Other material examined

MADAGASCAR: 1F,1Juv, same locality and data as holotype, general collecting at night (CASENT9009481).

Etymology

The specific name is a patronym for Mila Jane Macaulay in the hope that one day she will go to Andohahela to find this spider.

Diagnosis

Females of E.milajaneae sp. n., E.milloti sp. n., and E.lukemacaulayi sp. n. (and presumably E.pauliani, although the females are unknown) are distinguished from other “bourgini group” species by the enlargement of the U-shaped posterior bar of the internal genitalia. While the PB is present in other “bourgini group” species, in E.milajaneae sp. n., E.milloti sp. n., and E.lukemacaulayi sp. n., this bar is very large and curved into a u-shape (Figs 14B, 16B, 17B). In E.milajaneae sp. n. and E.milloti sp. n., each curved end of the PB extends at least to the middle of the “wings”. Eriaucheniusmilajaneae sp. n. is also distinguished by the rectangular shape of the posterior of the FSGP (Fig. 16B, arrow), and by the long neck that does not have a strong constriction, so that the “head” and “neck” are not clearly separated (Fig. 16A).

Etymology

The specific name is a patronym to honor Dr. Jacques Millot for his work describing Madagascan archaeids.

Diagnosis

Males distinguished from other “bourgini group” species except E.pauliani and E.lukemacaulayi sp. n. (and presumably E.milajaneae sp. n., although the males are unknown) by the presence of a lateral process on the conductor, which is narrow in E.lukemacaulayi sp. n. and E.milloti (Figs 14E, G–I, 17D–I, asterisk), but is broader in E.pauliani (Fig. 18C–D, F–H, asterisk). E.milloti sp. n. is distinguished from E.lukemacaulayi sp. n. by the quarter-moon-shape of the pedipalpal bulbs (Fig. 17D–F, J–L). Females of E.lukemacaulayi sp. n., E.milajaneae sp. n., and E.milloti sp. n. (and presumably E.pauliani) are distinguished from other “bourgini group” females by the enlarged, extended U-shaped PB (Figs 14B, 16B, 17B). The PB is smaller and thiner in other “bourgini group” females. In E.milloti sp. n. the curved ends of the PB extend all the way to the anterior edge of the FSGP (Fig. 17B), whereas in E.milajaneae sp. n. and E.lukemacaulayi sp. n. the curved ends do not go past the anterior of the “wings” (Figs 14B, 16B).

Description

Male holotype (USNMENT01377187, from Parc National Andohahela, Madagascar). Total length 2.18, carapace 0.91 long, 0.85 wide. Abdomen 1.23 long, 1.31 high. Carapace tilt angle 69.58°, tilt height (CtH) 1.89, constriction 0.47, head length 0.81, neck length 0.92. CtH divided by carapace length 2.08. Cephalon with AME on large bulge. Cephalon with 4 short post-ocular spines at the crown, not on protrusions. Chelicerae 1.81 long, and with short spine 0.30 from base of chelicerae and projecting downward (Fig. 17A). Femur I 2.53 long. Sternum 0.59 long, 0.38 wide. Carapace, chelicerae, and sternum reddish brown with white setae, although chelicerae lighter brown in middle. All legs with parts that are lighter brown and parts with darker annulations. Abdomen yellowish white, with scattered small and large dark brown areas with yellowish spots, and with tufts of white setae (Fig. 17A). Pedipalpal bulb with a membranous sac above the base of the embolus, with a greatly exposed broad and dark embolus that is encircled by the conductor (Fig. 17D–L). Conductor with a small narrow process (Fig. 17D–I, asterisk) followed by a constriction (Fig. 17H, arrow) that is then followed by a large bulge on the retrolateral side.

Type material

Other material examined

No other material examined.

Diagnosis

Male is distinguished from other “bourgini group” species by the distinctly shaped cephalon, that is triangular with a post-ocular pointed “head” (Fig. 18A), and also by the complete fusion of the sternum and carapace. Male is distinguished from other “bourgini group” species except E.milloti sp. n. and E.lukemacaulayi sp. n. (and presumably E.milajaneae sp. n., although the male is unknown) by the presence of a lateral process on the conductor (Figs 14G–I, 17D–I, 18F–G, asterisk), which in only E.pauliani is as wide as long.

Description

Male holotype (MNHN 13/1970, from Andohahelo, Fort Dauphin, Madagascar). Total length 2.39, carapace 0.97 long, 0.82 wide. Abdomen 1.30 long, 1.40 high. Carapace tilt angle 62.38°, tilt height (CtH) 2.57, constriction 0.54, head length 1.22, neck length 1.29 (Fig. 2). CtH divided by carapace length 2.65. Cephalon with AME on large bulge, and with postocular pointed head (Fig. 18A–B). Cephalon with 4 small post-ocular spines at the crown, not on protrusions. Chelicerae 2.07 long, and with a short spine 0.31 from base of chelicerae that projects perpendicular to the cheliceral cuticle. Femur I 2.99 long. Sternum 0.41 long, 0.65 wide, but completely fused to the carapace; measurements were taken based on approximations of where the original sternum edge would have been. Carapace and sternum reddish brown with white setae, chelicera light reddish brown. Legs beige with reddish brown annulations on tibia and metatarsus. Abdomen beige, mottled with brown and with tufts of white setae (Fig. 18A). Pedipalpal bulb with a membranous sac above the base of the embolus, with a greatly exposed embolus that is encircled by the conductor. Conductor with basal part an uneven ridge shape (see the basal “c” in Fig. 18F–G), with distal part that is narrow as it encircles the embolus, with a large, broad, bulge before tapering off. Conductor with lateral process that is as wide as long (Fig. 18F–G, asterisk). Embolus broad but tapering at tip to a long, fine wire (Fig. 18I); the embolus tip is broken off of the left pedipalp (Fig. 18G, arrow), but is entire in the right pedipalp (Fig. 18I).

Female: The female paratype at MNHN was not available for examination.

Variation

Natural history

Distribution

Known only from the type locality, Andohahelo, in southeast Madagascar (Fig. 32). The distribution point in Fig. 32 is an approximation since the holotype label does not have latitude/longitude data.

Comments

In E.pauliani the sternum is fused to the carapace, so the sternum measurements were based on an approximation of where the sternum border would be if this fusion was absent. In the type specimen the embolus tip is broken off in the left pedipalp (Fig. 18G – arrow), however the right pedipalp shows that the embolus tip is curved and much longer (Fig. 18I).

Diagnosis

Males and females are distinguished from other Eriauchenius, except E.mahariraensis and E.sama sp. n., by the presence of 4 small spines on the cephalon crown, and the cheliceral spine pointing perpendicular. Males of E.ratsirarsoni are distinguished from E.sama sp. n. and E.mahariraensis by lacking membranous tissue close to the conductor tip (Fig. 15D–L) and having a long conductor and conductor process that extends past the embolus tip (Fig. 19F, I). Females are distinguished by the presence of two lateral bulges on the FSGP and translucent “wings” (Fig. 19B, arrows).

Description

Male holotype (CASENT9012339, from Forêt Clasée Sandranantitra, Madagascar). Total length 1.67, carapace 0.80 long, 0.74 wide. Abdomen 0.81 long, 0.87 high. Carapace tilt angle 67.5°, tilt height (CtH) 1.35, constriction 0.46, head length 0.75, neck length 0.64. CtH divided by carapace length 1.69. Cephalon with AME on a very small bulge, and with 4 small post-ocular spines (anterior two are broken off, rudimentary or missing) on the crown of the cephalon, and 1 spine between the LE and median eyes (on each lateral side, for a total of 2; broken off on the right side). Chelicerae 1.34 long, and with a long spine 0.20 from base of chelicerae that projects perpendicular. Femur I 1.58 long. Sternum 0.55 long, 0.37 wide. Carapace, chelicerae, sternum, and legs reddish brown with white setae; patellas I–IV, legs II–III, tarsi and metatarsi I–IV pale yellow. Abdomen mostly dark brown with light circular patches, but with several light-yellow patches; abdomen with white and brown setae. Pedipalpal bulb with a membraneous sac above the base of the embolus, with a greatly exposed embolus that is encircled by the conductor (Fig. 19D–L). Conductor swirls around the broad, dark embolus and has a large, long process on the prolateral side (Fig. 19D–I, c1); conductor and process extend past the tip of the embolus (Fig. 19F, I).

Female paratype (CASENT9028378). Total length 1.89, carapace 0.86 long, 0.76 wide. Abdomen 0.87 long, 1.07 high. Carapace tilt angle 69.8°, tilt height (CtH) 1.44, constriction 0.54, head length 0.73, neck length 0.67. CtH divided by carapace length 1.68. Cephalon with AME on a small bulge, and with 4 small spines (anterior pair is rudimentary) on the crown of the cephalon, and missing spine between the LE and median eyes. Chelicerae 1.41 long, and with small spine 0.19 from base of chelicerae projecting perpendicular. Femur I 1.44 long. Sternum 0.56 long, 0.39 wide. Carapace and sternum orangeish brown; legs and chelicerae pale yellow. Abdomen mostly light yellow but with several dark brown areas with light circular patches; abdomen with white and brown setae. Female genitalia FSGP with two large lateral bulges (Fig. 19B, arrows), and nearly translucent “wings”, with PB present (Fig. 19B); one group of poreplates on each side of the bursa anterior; poreplates heavily sclerotized and raised, forming a bulge on the bursa (Fig. 19C).

Variation

Total length 1.67–1.70 (males; n=3), 1.70–2.47 (females; n=3); Carapace length 0.75–0.80 (males; n=3), 0.82–0.87 (females; n=3); Femur I 1.82–1.98 times the length of carapace in males (n=3), 1.63–1.75 in females (n=3); CtH divided by carapace length 1.65–1.69 in males (n=3), 1.68–1.70 in females (n=3). Average femur I length 1.47 in males (n=3) and 1.43 in females (n= 3). In all males the anterior pair of spines on the cephalon are either rudimentary or broken off; in females, one with anterior pair rudimentary, one with all 4 spines present, one with one anterior spine present and the other absent.

Natural history

In montane and primary rainforest from 450–1080 m in elevation, collected by sifting litter.

Distribution

Known only from central-eastern Madagascar (Fig. 31).

Nomenclatural remarks

One male specimen (CASENT9062876) has a pedipalpal bulb that is more heavily sclerotized than the other specimens, and also has a blunt conductor tip (Fig. 19K–L, arrows) rather than the tapering tips in the other two specimens (Fig. 19J, arrow). All three male specimens occur in close proximity so this is likely not a case of geographic variation. This specimen may be a new species and future molecular work as well as additional collection of specimens from more localities can resolve this issue.

Other material examined

Etymology

The specific name is a patronym to honor Dr. Michael Rix for his work describing Australian archaeids and examining their biogeographic patterns and evolutionary relationships.

Diagnosis

Males are distinguished from other “bourgini group” species by having a sharp process at the base of the pedipalpal tegulum (Fig. 20G–I, arrows). Females are distinguished from other “bourgini group” species by having the posterior bar curved towards the dorsal rather than the anterior (Fig. 20C), by having large broad “wings” and a narrow posterior elongation that is blunt at the end (Fig. 20C, arrow).

Other material examined

Diagnosis

Males and females are distinguished from other “bourgini group” species, except E.ratsirarsoni and E.mahariraensis, by having 4 spines on the cephalon, and the cheliceral spine pointing perpendicular. Males of E.sama sp. n. are also distinguished from other “bourgini group” species, by having a long process on the conductor that extends to the embolus tip (Fig. 21D–I, sclerite “c1”), and having the tip of the conductor point curving in a different direction than the more basal portion (Fig. 21G–I, arrow). Females are distinguished by the reduced FSGP, but the width divided by height is less than 2 (Fig. 21B).

Etymology

The specific name is a noun in apposition; ‘sama’ means ‘pelican’ in Malagasy.

Description

Male holotype (CASENT9012014, from Parc National Ranomafana, Madagascar). Total length 2.10, carapace 0.95 long, 0.85 wide. Abdomen 1.10 long, 1.26 high. Carapace tilt angle 72.5°, tilt height (CtH) 1.72, constriction 0.52, head length 0.82, neck length 0.81. CtH divided by carapace length 1.81. Cephalon with AME on a small bulge, and with 4 small post-ocular spines (although it is difficult to tell as the anterior pair is missing, rudimentary or broken off) on the crown of the cephalon, and 1 spine between the LE and AME (on each side, for a total of 2). Chelicerae 1.59 long, and with a spine 0.25 from base of chelicerae that projects perpendicular. Femur I 2.03 long. Sternum 0.63 long, 0.40 wide. Carapace, chelicerae, sternum, and legs reddish brown with white setae; patellas, tibias, and metatarsi light tan. Abdomen mottled with dark brown and light yellow patches, with light circular patches on the dark brown portions, with white setae. Pedipalpal bulb with a large membraneous sac above the base of the embolus, with a greatly exposed embolus that is encircled by the conductor (Fig. 21D–L). Conductor with a large bulge and a long process on the prolateral side that extends to the tip of the embolus (Fig. 21D–I, “c1” sclerite). In ventral view the left conductor curves in a clockwise direction, but the tip changes direction (Fig. 21H–I, arrow). Embolus broad and dark (Fig. 21G).

Female paratype (CASENT9012341). Total length 2.24, carapace 0.87 long, 0.80 wide. Abdomen 1.18 long, 1.48 high. Carapace tilt angle 67.5°, tilt height (CtH) 1.50, constriction 0.50, head length 0.78, neck length 0.73. CtH divided by carapace length 1.73. Cephalon with AME on a small bulge, and with 4 small post-ocular spines (one is broken off or missing) on the crown of the cephalon, and 1 spine between the LE and AME (on each side, for a total of 2). Chelicerae 1.54 long, and with small spine 0.29 from base of chelicerae. Femur I 1.82 long. Sternum 0.60 long, 0.40 wide. Cephalothorax colors as in male. Abdomen anterior light tan, and posterior dark brown with light circular patches; abdomen with white and brown setae. Female genitalia FSGP small and simple, with “wings” reduced (Fig. 21B); PB present; with poreplates in one group on each side of the bursa anterior (Fig. 21C).

Variation

no other known material.

Natural history

Female specimen was collected at 1700 m in elevation in forest by sifting litter.

Distribution

Known only from central-eastern Madagascar (Fig. 31).

Nomenclatural remarks: The male holotype and female paratype of E.sama sp. n. occur in different areas. The male and female were associated based on body size and carapace shape. Future molecular work as well as additional collection of specimens from more localities can help resolve this issue.

Etymology

The specific name is a noun in apposition; ‘zirafy’ means ‘giraffe’ in Malagasy.

Diagnosis

E.bourgini and E.zirafy sp. n. are distinguished from other “bourgini group” species by having two large protrusions on the crown of the cephalon (Fig. 22A). Females of E.bourgini and E.zirafy sp. n. are distinguished by the presence of two sclerotized invaginations on the bursa (Fig. 22C, arrows). Males of E.bourgini and E.zirafy sp. n. are distinguished by having a conductor with 4 processes in E.bourgini (Fig. 13D–K) and 5 in E.zirafy sp. n (Fig. 22D–K). Eriaucheniuszirafy sp. n. is distinguished from E.bourgini by lacking posterior extensions on coxae I (Fig. 13L), and by having a cluster of small bumps and pores on the conductor base of the male pedipalp (Fig. 22G–I,L, arrow).

Description

Male holotype (USNMENT01377199, from Parc National Masoala, Madagascar). Total length 1.79, carapace 0.75 long, 0.70 wide. Abdomen 0.97 long, 0.81 high. Carapace tilt angle 59.5°, tilt height (CtH) 1.72, constriction 0.32, head length 0.86, neck length 0.89 (Fig. 2). CtH divided by carapace length 2.29. Cephalon with AME on large bulge. Cephalon with 4 small post-ocular spines on the crown of the cephalon, with the posterior pair on large protrusions and the anterior pair not on protrusions, and 1 small spine between the LE and AME (on each side, for a total of 2). Chelicerae 1.50 long, and with a short spine 0.68 from base of chelicerae, projecting downward. Femur I 1.89 long. Sternum 0.50 long, 0.32 wide. Carapace, chelicerae, and sternum dark reddish brown with white setae. Coxae and legs lighter brown, with darker annulations on tibiae and metatarsi. Abdomen mottled brown and beige, with a bright white patch on each lateral side, with tufts of white setae (Fig. 22A). Pedipalpal bulb with a small membranous sac above the embolus base, with the base of the conductor small and triangular with a cluster of small bumps and pores (Fig. 22G–I,L, arrow), and with the remainder of the conductor with 4 long processes as it wraps around the embolus, although one process is bifurcating (Fig. 22D,G–I, c3 and c4) for a total of five processes (Fig. 22D–I, c1–c5). The embolus is thick, contains two processes, and has membranous parts.

Variation

Total length 1.79–1.80 (males; n=3); Carapace length 0.72–0.75 (males; n=3); Femur I 2.54–2.72 times the length of carapace in males (n=3) and 2.54 times the length of the carapace in females (n=1). CtH divided by carapace length 2.29–2.44 in males (n=3). Average femur I length 1.96 (males; n=3).

Natural history

Specimens were collected from 195–700 m in elevation in rainforest by beating vegetation and general collecting.

Distribution

Known only from areas around Parc National Masoala in northeastern Madagascar (Fig. 31).

GenusMadagascarchaeagen. n.

Type species

Archaeagracilicollis Millot, 1948

Etymology

The name refers to the Madagascan distribution and is feminine in gender.

Diagnosis

Distinguished from all other archaeids by the presence of six protrusions, each with a small spine, on the crown of the cephalon, and by the presence of a retrolateral apophysis on the male pedipalpal patella.

Distribution

Madagascar

Discussion

We elevate the “gracilicollis group” (Wood 2008) to genus level based on phylogenetic analysis that strongly supports two monophyletic clades, Eriauchenius and Madagascarchaea gen. n., that have diversified on Madagascar (Fig. 1) (Wood et al. 2015; Wood et al. 2012). These two genera are not sister clades, but instead Afrarchaea is sister to Eriauchenius in Wood et al (2015), and Afrarchaea is sister to Madagascarchaea gen. n. in Wood et al (2012). Eriauchenius and Afrarchaea are never recovered as sister clades.

Madagascarchaea gen. n. contains two main clades, the “vadoni group” and the “jeanneli group” (Fig. 1). The “vadoni group” contains M.borimontsina, M.legendrei, M.vadoni, M.fohy sp. n., and M.rabesahala sp. n., and these species share the following traits: a retrolateral apophysis on the male pedipalpal femur (compare fig. 12A with fig. 12B in Wood 2008) and a highly reduced FSGP that lacks “wings” (Figs 23B, 24B, 25B). The female genitalia of “vadoni group” species do not have interspecific variation. Instead, somatic traits are more useful for identifying “vadoni group” species. We split what was previously considered M.legendrei into two species: M.legendrei and M.rabesahala sp. n. For clarity, we describe both M.legendrei and M.rabesahala sp. n. here. In addition, there is a single female specimen (CASENT9046596), denoted as “Madagascarchaea sp. 1” in Fig. 1 that may be a new species, however, since this is the only known specimen and females are difficult to diagnose, we chose not to describe it (collection information: Madagascar, Fianarantsoa, Réserve Forestiére d’Agnalazaha, Mahabo, 42.9 km 215° SW Farafangana, 23°11'38"S, 47°43'23"E, 20 m, 19 Apr 2006, littoral rainforest, maxi winkler litter extraction, B.L. Fisher et al.).

M.jeanneli, M.ambre, M.lotzi sp. n., and M.moramora sp. n. belong to the “jeanneli group” and are part of a species complex distinguished from other Madagascarchaea by having an elongated and pointy head (Figs 26A, 27A, 28A), a triangular abdomen that is either straight across the posterior (Fig. 28A) or invaginated (Figs 26A, 27A), and a conductor that is a large triangular piece in the prolateral view (Figs 26D, 27D). The current study splits the M.jeanneli of Wood (2008) into several different species: M.jeanneli, M.lotzi sp. n., and M.moramora sp. n. For clarity, we also redescribe M.jeanneli here. M.ambre, is considered a part of the “jeanneli-complex” but is not redescribed here, see Wood (2008) for a complete description. These four species form a monophyletic group (Fig. 1), although M.jeanneli is not included in the phylogeny. Females cannot be distinguished. For the males, embolus shape is the main morphological difference among species.

Madagascarchaea gen. n.: to supplement “Gracilicollis group” key

When using the identification key from Wood (2008), if a specimen is identified as either M.jeanneli or M.ambre, then use the following key to separate M.jeanneli, M.ambre, M.lotzi sp. n., and M.moramora sp. n.; females of these four species are indistinguishable:

Embolus with a deeper bifurcation, both parts of bifurcation different thickness (Figs 26F, I, L, 28E–F, compare shape of “a” with shape of “p”); abdomen invaginated or straight across the posterior in both sexes (Fig. 28A, arrow)

2

2

Anterior portion of embolus bifurcation thinner than the posterior part, and jutting out past the conductor in the ventral view (see fig. 21 in Wood 2008); usually abdomen straight across the posterior in both sexes (as in Fig. 28A, arrow; also see fig. 1C in Wood 2008)

Anterior portion of embolus bifurcation thick and blunt at tip (Fig. 28D–F, “a”), posterior portion with a bifurcation at tip with each piece of unequal width (Fig. 28D–F, “p”); posterior part of abdomen usually straight in both sexes (Fig. 28A, arrow)

M.moramora

–

Anterior portion of embolus bifurcation tapering off to a point (Fig. 26D–L, “a”), posterior portion with a bifurcation at the tip (Fig. 26E–F, H–I); posterior part of abdomen usually invaginated in both sexes (Fig. 26A)

M.jeanneli

When using the identification key from Wood (2008), if a specimen is identified as M.legendrei, then use the following key to separate species of M.legendrei, M.fohy sp. n., and M.rabesahala sp. n.:

1

In males and females, cephalon crown rounded (Figs 23A, 24A)

2

–

Cephalon crown not as round in both sexes (Fig. 25A)

M.rabesahala

2

In males, pedipalpal bulb, with a sclerotized rod-shaped piece on embolus (Fig. 23H, J, arrow); presence of a cymbium process that has setae on the retrolateral side of the cymbium (Fig. 23C); females of M.fohy and M.legendrei are indistinguishable

Etymology

The specific name is a noun in apposition; ‘fohy’ means ‘shorty’ in Malagasy.

Diagnosis

Distinguished from all other archaeids, except M.vadoni, M.legendrei, M.rabesahala sp. n., and M.borimontsina (likely, because the male is unknown) by having a retrolateral apophysis on the distal side of both the palpal femur and patella (fig. 12B from Wood 2008), and because the FSGP is highly reduced and lacks “wings” (Figs 23B, 24B, 25B). M.fohy sp. n. is distinguished from M.vadoni, M.legendrei, and M.rabesahala sp. n. by having a heavily sclerotized rod-shaped sclerite on the palpal bulb (Fig. 23H, J, arrow), and by the presence of a retrolateral protrusion on the cymbium that has setae (Fig. 23C, arrow). M.fohy sp. n. is further distinguished from M.vadoni by having 6 spines that are not on protrusions on the cephalic crown, opposed to the small protrusions seen in M.vadoni (Wood, 2008: fig. 18A, C); from M.borimontsina by having less than 6 true teeth on the cheliceral retromargin (Wood 2008), by lacking a point on the AME bulge (Wood 2008: fig. 18D), and by lacking a cheliceral swelling on the posterior-basal side of the chelicerae (Wood 2008: fig. 18D).

Variation

Total length 1.67–1.97 (females; n=2); Carapace length 0.71–0.75 (females; n=2); Left and right femur I missing in one female (CASENT9015493). Femur I 2.73 times the length of carapace in females (n=1); CtH divided by carapace length 1.92–2.07 in females (n=2).

Natural history

Specimens were collected in disturbed and undisturbed rainforest from 26–950 m in elevation by general collecting.

Distribution

Known only from northeastern and central-eastern Madagascar (Fig. 33).

Diagnosis

Distinguished from all other archaeids, except M.vadoni, M.rabesahala sp. n., M.fohy sp. n., and M.borimontsina (presumably, because the male is unknown) by having a retrolateral apophysis on the distal side of the male pedipalpal femur and patella (see fig. 12B in Wood 2008), and by the shape of the FSGP that is highly reduced and lacks “wings” (Fig. 24B). M.legendrei is distinguished from M.vadoni by having a rounded cephalon with the cephalic spines not on protrusions (Fig. 24A), rather than the larger protrusions seen in M.vadoni (compare figs 18A and 18C in Wood 2008); from M.borimontsina by having less than 6 teeth on the cheliceral retromargin, by having a rounded bulge that the AME rests upon, and by lacking a bulge on the posterio-basal side of the chelicerae (see fig. 18D in Wood 2008); from M.fohy sp. n. by lacking a heavily sclerotized rod-shaped sclerite on the palpal bulb (Fig. 23H,J, arrow), and by lacking setae on the retrolateral cymbial protrusion (Figs 23C, 24C, arrow); and from M.rabesahala sp. n. by having a more rounded cephalon, and by having a comparatively smaller and less sclerotized triangular basal portion of the conductor (compare basal “c” in Figs 24G, 25G).

Description

Male (based on CASENT9012333, from Parc National Ranomafana, Madagascar). Total length 1.67, carapace 0.74 long, 0.58 wide. Abdomen 0.90 long, 1.13 high. Carapace tilt angle 61.9°, tilt height (CtH) 1.42, constriction 0.28, head length 0.58, neck length 0.84. CtH divided by carapace length 1.92. Cephalon with AME on large bulge, and with 6 short post-ocular spines at the crown, not on protrusions, and 1 short spine between the LE and AME (on each side, for a total of 2; see fig. 18A in Wood 2008). Chelicerae 1.53 long, and with a small spine 0.34 from base of chelicerae and projecting downward (Fig. 24A). Femur I 2.35 long. Sternum 0.49 long, 0.29 wide. Carapace, chelicerae, sternum and legs reddish brown with white setae. Legs with darker annulations on tibiae and metatarsi. Abdomen dark brown with lighter circular patches throughout, with white setae (Fig. 24A). Pedipalpal bulb of the “vadoni group” form (Fig. 24D–I): pedipalpal bulbs elongated with conductor swirling around a mostly membraneous embolus; conductor base triangular (Fig. 24D,G), with remainder of conductor elongate and cradling embolus, and with a curved tip; MA translucent and fans out (Fig. 24E–F, H–I). Cymbium with small retrolateral protrusion that lacks setae (Fig. 24C, arrow).

Etymology

The specific name is a noun in apposition and commemorates Gisèle Rabesahala, a Malagasy activist and politician.

Diagnosis

Distinguished from all other archaeids, except M.vadoni, M.legendrei, M.fohy sp. n., and M.borimontsina (presumably, because the male is unknown) by having a retrolateral apophysis on the distal side of the male pedipalpal femur and patella (see fig. 12B in Wood 2008), and because the FSGP is highly reduced and lacks “wings” (Fig. 25B). M.rabesahala is distinguished from M.vadoni by having the cephalic spines not on protrusions (Fig. 24A), rather than the protrusions seen in M.vadoni (see fig. 18C in Wood 2008); from M.borimontsina by having less than 6 teeth on the cheliceral retromargin, by having a rounded bulge that the AME rests upon, and by lacking a bulge on the posterio-basal side of the chelicerae (see fig. 18D in Wood 2008); from M.fohy sp. n. by lacking a heavily sclerotized rod-shaped sclerite on the palpal bulb (Fig. 23H,J, arrow), and by lacking a retrolateral cymbial protrusion (Fig. 25C, arrow); and from M.legendrei by having a cephalon that is not as perfectly rounded (compare cephalon shape in Figs 24A, 25A), and by having a larger and more sclerotized triangular basal portion of the conductor (compare basal “c” in Figs 24G, 25G).

Variation

Total length 1.56–1.66 (males; n=5), 1.91–2.07 (females; n=4); Carapace length 0.72–0.75 (males; n=5), 0.74–0.81 (females; n=4); Femur I 2.69–2.86 times the length of carapace in males (n=5), 2.38–2.69 in females (n=4); CtH divided by carapace length 1.93–2.12 in males (n=5), 1.94–2.15 in females (n=4); Average femur I length 2.03 in males (n=5), 1.96 in females (n= 4). The spine between the LE and AME is present is 3 males out of 5, and 4 females out of 4.

Natural history

Specimens were collected in montane rainforest from 960–1300 m in elevation by beating low vegetation, by sifting litter, by beating and sweeping, and by general collecting.

Distribution

Known only from central eastern Madagascar (Fig. 33).

The new ‘pointy-head’ “vadoni group” species and a redescription of M.jeanneli

Diagnosis

Distinguished from all Madagascarchaea, except other ‘pointy head’ species M.ambre, M.lotzi, and M.moramora sp. n., by having a conductor that is a concave triangular shape (Fig. 26D–E, J–H, J–K), and by having an abdomen that is invaginated across the back (Fig. 26A), rather than rounded (as in Fig. 23A). Typically M.jeanneli can be distinguished from M.ambre and M.moramora sp. n., by having an abdomen that is invaginated in the posterior (Fig. 26A), rather than straight (Fig. 28A). M.jeanneli is further distinguished from M.moramora by having the anterior portion of the embolus taper off to a point (Fig. 26G–I, “a”) rather than being broad and blunt (Fig. 28D–F, “a”), and from M.ambre by having the anterior piece of the embolus broad and curved (Fig. 26G–I, “a”) and not straight, narrow, and jutting out past the conductor in the retrolateral direction (see fig. 21 in Wood 2008). M.jeanneli is distinguished from M.lotzi sp. n. by having an embolus with a very deep bifurcation (Fig. 26I, L), and with the posterior portion of the embolus having an additional bifurcation at the tip (Fig. 26F, I, “p”).

Natural history

Specimens were collected in montane rainforest from 960–1300 m in elevation by beating low foliage, low vegetation, by beating and sweeping, by general collecting at night, and by hand collecting at night in vegetation.

Distribution

Known only from central eastern Madagascar (Fig. 34).

Nomenclature remarks

Previous work (Wood 2008) lumped into M.jeanneli what we are here calling several different species. The M.jeanneli syntypes include two females, and females of the “jeanneli complex “(M.ambre, M.jeanneli, M.lotzi sp. n., and M.moramora sp. n.) cannot be confidently distinguished. However, the type locality of “La Mandraka” is an area about 40 kilometers east of Antananarivo, and so we propose that the female syntypes are conspecific to the males found in this area. The male and female specimens that were described and illustrated as M.jeanneli in Wood 2008 are here being called M.lotzi n. sp.

Etymology

The specific name is a patronym to honor Dr. Leon Lotz for his work in describing the South African and Madagascan archaeids.

Diagnosis

Typically M.lotzi sp. n. can be distinguished from the northern species, M.ambre and M.moramora sp. n., by the presence of an invagination in the abdomen posteriorly (Fig. 27A, arrow). M.lotzi sp. n. is further distinguished from M.moramora sp. n. by having a bifurcation in the embolus that is more shallow (Fig. 27E–F, H–I) rather than the deeper bifurcation of M.moramora sp. n. (Fig. 28E, F), and from M.ambre by having the anterior portion of the embolus not narrow and jutting out past the conductor in the retrolateral direction (see fig. 21 in Wood 2008). M.lotzi sp. n. is distinguished from M.jeanneli by having an embolus with a more shallow bifurcation, and with the posterior portion of the embolus lacking a large bifurcation at the tip (Fig. 27E–F, H–I).

Natural history

Specimens were collected in montane rainforest from 900–1200 m in elevation by general collecting, beating and puffing, general collecting at night, beating vegetation, and beating clumps of dead, dry foliage.

Distribution

Known only from Parc National Ranomafana in central eastern Madagascar (Fig. 34).

Other material examined

Etymology

The specific name is a noun in apposition; ‘mora mora’ means ‘easy easy’ in Malagasy.

Diagnosis

M.moramora sp. n. can be distinguished from the southern species, M.jeanneli and M.lotzi sp. n., by the abdomen being straight across the posterior (Fig. 28A, arrow) rather than invaginated (Figs 26A, 27A). M.moramora sp. n. is further distinguished from M.lotzi sp. n. by having a deep embolus bifurcation (Fig. 28E–F) compared to the shallow bifurcation of M.lotzi sp. n. (Fig. 27E–F, H–I), and from M.ambre by having the anterior portion of the embolus bifurcation not narrow and jutting out past the conductor in the retrolateral direction (see fig. 21 in Wood 2008). M.moramora sp. n. is distinguished from M.jeanneli by having the anterior portion of the embolus bifurcation broad and blunt (Fig. 28F), rather than tapering (Fig. 26F, I), and having the posterior portion with a bifurcation at the tip with one side narrower than the other (Fig. 28D–E), whereas in M.jeanneli the bifurcation at the tip is equal sized on each side (Fig. 26F, I).

Description

Male holotype (USNMENT01377197, from Mikira Forest, Madagascar). Total length 1.53, carapace 0.67 long, 0.46 wide. Abdomen 0.83 long, 0.92 high. Carapace tilt angle 57.2°, tilt height (CtH) 1.25, constriction 0.35, head length 0.73, neck length 0.63. CtH divided by carapace length 1.86. Cephalon with AME on large bulge, and with 6 short post-occular spines at the apex, not on protrusions, and 1 short spine between and posterior to the LE and median eyes (see fig. 18B in Wood 2008). Chelicerae 1.21 long, and with a small spine 0.28 from base of chelicerae and projecting downward (Fig. 28A). Femur I 1.86 long. Sternum 0.44 long, 0.24 wide. Carapace, chelicerae, and sternum reddish brown with white setae. Legs light brown with darker annulations throughout. Abdomen mottled with dark brown areas and lighter whitish areas, anterior of abdomen with large white patch, abdomen interspersed with white and brown setae (Fig. 28A). Posterior edge of abdomen straight and not invaginated (Fig. 28A, arrow). Conductor concave and triangular; MA dark, thick, and curves anteriorad; S1 present as a thin ridge; Embolus dark, with a deep bifurcation, with anterior portion broad and blunt at the tip, and with posterior portion with bifurcation at the tip, of which both sides are curved but one side is more narrow (Fig. 28D–F).

Female paratype (USNMENT01377198). Total length 1.98, carapace 0.79 long, 0.52 wide. Abdomen 1.09 long, 1.35 high. Carapace tilt angle 48.2°, tilt height (CtH) 1.48, constriction 0.35, head length 0.82, neck length 0.70. CtH divided by carapace length 1.87. Cephalon as in male. Chelicerae 1.42 long, and with a short spine 0.38 from base of chelicerae and projecting downward. Femur I 12.25 long. Sternum 0.49 long, 0.26 wide. Colors as in male. Posterior edge of abdomen straight and not invaginated. Genitalic bursa divided down middle by a sclerotized piece on the anterior-ventral side, with several groups of poreplates on either side; FSGP with two strong points arising from either side of anterior edge, having ‘wings’, and lacking posterior elongation (Fig. 28B–C).

GenusAfrarchaeaForster & Platnick, 1984

Type species

Diagnosis

Distinguished from all extant genera by the presence of a prominent keel on the FSGP (see fig. 58 of Forster and Platnick (1984), fig. 3 of Lotz (2006), and figs 1c–d and 6b–c of Lotz (1996)). The keel likely has been lost in A.royalensis and A.ngomensis given the central phylogenetic placement of these two species (Fig. 1).

Distribution

South Africa.

Discussion

Phylogenetic analysis based on molecular data recovered a monophylectic Afrarchaea with strong branch support (Wood et al. 2015). This research supports the transfer of A.cornutus from Eriauchenius to Afrarchaea and the transfer of E.mahariraensis from Afrarchaea to Eriauchenius (Fig. 1). Furthermore, females of A.cornutus have a FSGP keel typical of Afrarchaea (see fig. 3 of Lotz 2006) and females of E.mahariraensis do not (Fig. 15B–C).

Regarding Afrarchaea distribution, Legendre (1970) reported on a male and female specimen (not examined for this study) collected at Manjakatompo, Madagascar, which he identified as Afrarchaeagodfreyi. However, Legendre noted that these specimens were different from A.godfreyi in some details (i.e., the absence of prominent cephalic spines and the absence of abdominal sclerotization). These differences suggest that these specimens are not A.godfreyi. Unfortunately, in the CAS collections only a single juvenile specimen has been collected from Manjakatompo. This specimen also has reduced spines on the cephalon (only one rudimentary spine is present), which is again suggestive that this specimen is not A.godfreyi. Furthermore, many Eriauchenius species endemic to Madagascar have the short fat “neck” typical of most Afrarchaea (other than A.cornutus and A.ansieae) and also have reduced spination on the cephalon. The species E.fisheri, E.goodmani sp. n., E.harveyi sp. n., E.mahariraensis, E.ratsirarsoni, E.sama sp. n., and E.wunderlichi sp. n. all superfically resemble Afrarchaea in terms of carapace shape (Figs 9A, 10A, 11A, 12A, 15A). E.mahariraensis and E.fisheri were originally described as Afrarchaea based solely on carapace shape (Lotz 2003). Current research on archaeids shows that carapace shape has evolved in parallel, with similar morphs evolving repeatedly (Wood 2017; Wood et al. 2015; Wood et al. 2007), suggesting that carapace shape is not a good diagnostic trait for these genera. Phylogenetic analysis (Wood et al. 2015) and morphological examination of numerous African and Madagascan specimens suggests instead that the African and Madagascan species are short-range endemics that are restricted to either southern Africa or Madagascar, but not both (Fig. 1). Furthermore, after over 10 years of extensive collecting in Madagascar by CAS researchers, specimens of Afrarchaeagodfreyi have never been found. For these reasons we propose that the distribution of Afrarchaeagodfreyi be restricted to South Africa until evidence suggests otherwise.

Figure 29.

Distribution map for Eriauchenius species.

Figure 30.

Distribution map for Eriauchenius species.

Figure 31.

Distribution map for Eriauchenius species.

Figure 32.

Distribution map for Eriauchenius species. ** The distribution for E.pauliani is an approximation as the holotype does not have latitude/longitude data.

Figure 33.

Distribution map for Madagascarchaea species.

Figure 34.

Distribution map for Madagascarchaea species.

Acknowledgements

This study was funded through a grant from the Danish National Research Foundation to the Center for Macroecology, Evolution, and Climate (DNRF96), and through a National Science Foundation Postdoctoral Fellowship (1202873). Access to and loan of specimens was made possible by Charles Griswold, Lauren Esposito and Darrel Ubick at CAS, Norman Platnick at AMNH, Petra Sierwald at FM, Christine Rollard at MNHN, Janet Beccaloni at BMNH, and Jonathan Coddington at USNM. Thanks to the Fisher/Griswold Arthropod Survey team, who were responsible for collecting and processing most of the specimens studied. Thanks to Benjamin Andriamihaja, Directeur Général of the Madagascar Institut pour la Conservation des Ecosystèmes Tropicaux (MICET) for help with logistics and Harin’Hala Hasinjaka (Rin’ha) of that institute and Balsama Rajemison for arranging collecting permits. Thanks to Michael Rix and Leon Lotz for reviewing and commenting on this manuscript.

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